Feddes Repertorium 103 (1992) 5 - 6, 357 - 361 Berlin, August 1992

Sardar Patel University, Faculty of Science, Department of Biosciences. Vallabh Vidyanagar, Gujarat, India

V. THOMAS

Structure and biology of floral nectary in Allamanda cathartica L.

(Apocynaceae)

With one Plate

Summary Zusammenfassung
Structure and biology of floral nectaries in Allumunda Struktur und Biologie der Bliitennektarien von Allu-
were studied. Disc nectary present around the ovary mandu werden untersucht. Rund um den Frucht-
was found to be five lobed, and differentiated into knoten sind die Nektarien flach und funfgeteilt. Sie
epidermis, secretory zone and sub secretory zone. sind unterteilt in Epidermis, Sekretionszone und den
Nectary traversed by vasculature, possesses both Bereich unterhalb der Sekretionszone. Die Nektarien,
xylem and phloem. The nectar is coming out through die von GefaBen durchzogen werden, haben sowohl
stomata and it contains an equal concentration of Xylem als auch Phloem. Der Nektar tritt durch die
glucose and fructose. The major visitors to suck nectar Stomata aus und enthalt in gleicher Konzentration
arc wasp and ants. Glukose und Fruktose. Die haufigsten Besucher, um
den Nektar zu genieBen, sind Wespen und Ameisen.
Key w o r d s : Allamanda catharticu; floral nectary;
necrosis; vasculature.

Introduction in the nectary of Pedilanthus (DAVEand PATEL

1975). According to RAO and GANGULI (1963)
Family Apocynaceae is highly neglected for the the nectaries in Apocynaceae are carpelloid.
study of nectaries except one deed in Vinca by Occurrence of carpelloid nectary in this family
RACHMILEVITZ and FAHN(1973) and ZALAet has been reported by FALLEN (1983, 1986) in
al. (1976). PATELand INAMDAR (1972) have Anechites, Funturnia, Kopsia, Mandeecilla, Pre-
made an extensive survey for extrafloral necta- stonia and Voacanga. The present study in-
ries (EFN) in Gentianales but they have not corporates the structure and biology of floral
reported the occurrence of EFN in any of the nectaries in Allamanda cathartica L.
members of Apocynaceae. There are certain
reports about the occurrence of EFN in this Materials and methods
family viz., Vinca (SCHLEE1977) Allamanda
(ELIAS 1983), Holarrhena (INAMDAR et al. 1985), Nectary samples of Allamundu catharticu L. were
Plumeria (MOHAN and INAMDAR1986) and collected from Sardar Patel University Botanical
Nerium (JAINet al. 1986). But later on the basis Garden, and fixed in F.A.A. (JOHANSON 1940). Fixed
of histochemistry and exudate analysis. could materials were dehydrated through graded series of
Tertiary Butyl Alcohol (SASS1958) and embedded in
recognize that the secretory structures present
'Tissue prep' of 56.5 " C . Serial sections of 6 - 8 pm
in the above mentioned genera are colleters, were cut on Spencer 820 rotary microtome and stained
glands secreting mucilage or resinous material, with Safranin-Fast green for general histology. The
rather than E F N (THOMAS1991). The colleters following histochemical stains such as PAS (JENSEN
have been mistaken for nectaries, due to its 1962), CBB (EKLAVYA1979) and Oil red 0 (LILLIE
superficial identity, distribution pattern etc. as and FULLMER 1976) used for starch, protein and lipid

24 Feddcs Repsrt. 103 (1992) 5 - 6

358 Feddes Repert., Berlin 103 (1992) 5-6

Fig. 1
A - disc nectary (arrow). x 3 ; B - nectary showing a constriction (arrow) on the adaxial side. x 95:
C - branched vasculature (arrows) in the nectary. x 80; D - transection of nectary showing stomata and
laticifer, x 80; E - a hole on the lower portion of the corolla tube, x 8

respectively. Thin layer chromatography (TLC) of Results

the nectar was carried out for sugars and aminoacids.
TLC was carried out on silica gel plates using solvents
n-butanol, acetic acid water: 4, 1, 5, v/v; and 8, 8, 2, Allumandu cathurtica have 5 lobed, pale yellow
v/v for sugars and aminoacids respectively. Aniline coloured disc nectary, in which about half of
diphenyl amino reagent was sprayed for the detection the superior ovary is sunken (Fig. I , A). Ovary
of sugars, and the ninhydrin test (PLUMMER 1982) in an open flower appears morphologically as
was performed for aminoacids. Quantitatively sugars
syncarpous due to a postgenital carpel fusion
were estimated by anthrone method and Somogyi
Nelson Method (PLUMMER 1971). Observation and to the free carpels (FALLEN 1985). Transection
photomicrographs were taken by Zeiss Photomi- of the nectary through the ovary part shows a
croscope. constriction on the adaxial side of the nectary,
V. THOMAS,
Floral nectary in Allumandu 359
exactly opposite to the region where the carpels Discussion
fused postgenitally (Fig. I , B).
In transection nectary is 30 - 35 layered In Apocynaceae the nectaries or disc lobes
thick, constituted of loosely arranged 'thin frequently are separate, and 2-5 in number,
walled cells. Nectary is differentiated into but occassionally they cohere to form a nearly
three zones, viz., epidermis, secretory zone uniform, annular structure (WOODSONand
and sub-secretory zone. A row of radially MOORE 1938). Nectaries contain definite
elongated epithelial cells constitute the epi- vascular tissue associated with that of gynoe-
dermis, covered externally by a thin cuticle. cium proper indicates that the nectaries definite-
Stomata are present on the nectary surface ly are appendages and not merely meaningless
(Fig. 1,D). During the development paren- hypertrophies of the receptacle. The traces of
chymatous nectariferous tissue is compactly the vascular bundles from and to the nectaries
arranged, without much intercellular spaces, provide a valuable basis for explaining the
but towards maturity the secretory cells become origin of nectaries (AGTHE1951; FREY-WYSS-
less compact, giving rise to prominent inter- LING 1955). After a study of the floral anatomy
cellular spaces. Nectariferous tissue in the devel- of Apocynum WOODSON(1930) concluded that
oping stages show abundant starch grains but the nectaries of Apocynaceae should be inter-
during secretion they are broken down. Protein preted as carpellodes. WOODSONand MOORE
and lipid are present sparingly in the secretory (1938) also supported the above after a study
stage. A characteristic feature of the nectary is of numerous additional genera, particularly the
the presence of vasculature (Fig. 1, C) and non- polycarpellary species of Plriocarpa murica. The
articulated latificer (Fig. 1, D). Vasculature in present author also agrees to the above state-
the nectariferous tissue is profusely branched ment because of the presence of vasculature to
and have connection with the floral trace. the nectary and the constriction found on the
The nectaries are functional only for a short adaxial side of the nectary, which may be a part
period of time. Nectar comes out through the of gynoecial development in Allarnanda.
stomata and accumulates in the space between The floral biology and pollination mecha-
the ovary and corolla tube. Accumulation of nism of various members of Apocynaceae have
nectar can be seen in the corolla tube in the obtained considerable attention from the bota-
morning hours of the day on which the flower nists (SPRENGEL 1793; DELPINO 1867; RAOand
opens. Analysis of the nectar shows the presence GANGULI1963; SHICK1980, 1982; FALLEN
of both sugars and aminoacids. The glucose- 1986). In the present study it is found that the
fructose and sucrose ratio in the nectar is almost floral nectaries in Allaniancla is definitely in-
equal. volved in pollination by attracting pollinators,
The major visitors to suck nectar from Alla- providing the floral reward - nectar in return.
manda flower are wasp and ants. The wasp visits Nectar robbing is a behaviour exhibited by
first and make a hole (Fig. 1, E) at the base of some species of birds, bees and ants in which
the corolla tube to suck the nectar, in the early nectar is obtained by them through holes in the
morning hours. Later, ants enter into the corolla corolla tube (INOUYE 1983).
tube for sucking the nectar through the hole LOKEN(1949) subdivided the nectar robbers
which has been made by the wasp. The sagittate into primary nectar robbers, or those species or
anther obstruct the entry of visitors into the individuals which make the holes and then
corolla tube through the mouth of the corolla. extract the nectar, and secondary rubbers, or
Corolla remains attached to the flower for those species or individuals which obtain nectar
one or two days after the flower opening. by using holes made by primary robbers.
Meanwhile nectary becomes functionless and Amongst the nectar robbers in Allarnanda, the
turns its colour slowly into pale brown. This is wasp is the primary robber and the ants are the
the visible sign of a senescent nectary. In a secondary robbers. Contrarily BENTLEYand
senescent nectary the cells are highly vacuolated ELIAS(1983) described that injured flowers of
and have lost, its shape. Dried nectariferous some tropical species of Apocynaceae produce
tissue can be seen as a rind at the base of the a sap which is likely to be a chemical deterrent
capsular fruit. to nectar robbing. The present author also

24'
360 Feddes Repert., Berlin 103 ( 1 992) 5 - 6

noticed holes on the corolla tube of Tubebuia lanrlius tithymaloides, Euphorbiaceae). Amer. J.
and Thunbergia for nectar robbing. Bot., 62, 808-812.
Based on the quantitative relationships be- DELPINO, F., 1867, Sugli apparecchi della feconda-
zione nelle plante antecarpee (Fanerogame). Som-
tween sucrose and glucose-fructose, three main
mario di osservazione fatte negli anni 1865 - 1866.
types of nectar have been distinguished: (1) Firenze.
sucrose dominant, (2) glucose-fructose domi- DELPINO, -
F., 1868 1875, Ulteriori, osservazioni sul-
nant, (3) those with an equal ratio of sucrose: la dicogamia nel regno Vegetale. I & 11. Atti Soc.
glucose-fructose (FAHN1949; PERCIVAL 1961; Italiana Sci. Natur., 11, 12.
BAHADURet al. 1986). They also suggested that . , A technique for making CBB
E K L A V Y A , ~1979,
in general nectar from concealed nectaries are stained sections of paraffin and resin-embedded
dominacd by sucrose, where as comparatively tissue. Indian J. Bot., 2, 73-75.
open flowers have more of reducing sugars. In ELIAS.T., 1983, Extrafloral nectaries: Their structure
the present study it is found that glucose- and distribution. In: The biology of nectariesied.
fructose concentration is equal in Allamunda by D. BENTLEY; T. ELIAS.New York.
nectar. FAHN,A., 1949a, Studies in the ecology o f nectar
secretion. Palestine J. Bot., Ser. 4. 207-224.
Disc nectary have been reported in many
FAHN,A.. 1949b, Secretory tissues in plants. London.
families by FAHN(1979). Both morphologically
FALLEN, M. E., 1983, A systematic revision of Ant,-
and anatomically the nectary of Allamanda is chiles (Apocynaceae). Brittonia, 35, 222 - 231,
identical to those in Bignoniaceae (see SUB- FALLEN, M. E., 1985, The gynoecial development and
RAMANIAN and INAMDAR 1985; THOMAS and systematic position of Allamunda (Apocynaceae).
DAVE1991), with the presence of stomata and Amer. J. Bot., 72, 572-579.
inter-cellular spaces - a characteristic feature FALLEN, M. E., 1986, Floral structure in the Apocy-
of many floral nectaries (WADDLE and LERSTEN naceae: Morphological, functional and evolutio-
1973; FAHN 1979). Nectary have a cuticular nary aspects. Bot. Jahrb. Syst., 106, 245-286.
insulation, which is considered as an adoptation FREY-WYSSLINC, A., 1905. The phloem supply to the
against transpiration loss (THOMAS and DAVE nectaries. Acta Bot. Need., 4. 358-369.
1991). The vasculature to the nectaries is well INAMDAR,J. A.; MOHAN,J . S. S.; SUBRAMANIAN,
R. B., 1985, Extrafloral nectaries of Holurrhenu
developed, profusely branched and phloem do-
anridysenrericn (L.) WALL.ex G. DON. Develop-
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vasculature in a structure is directly proportio- Res., 137 - 148.
nal to its size and it is not necessarily related INOUYE, D. W., 1983, The ecology of nectar robbing.
to any state of advancement. The probable In: The biology of nectaries/ed. by D. BENTLEY;
physiological function of laticifers in the nectary T. ELIAS.New York.
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